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A Study of Right and Left Atrial Receptors 596 J. Physiol. (I953) 120, 596-6I0 A STUDY OF RIGHT AND LEFT ATRIAL RECEPTORS By A. S. PAINTAL* From the Department of Physiology, University of Edinburgh (Received 1 December 1952) The satisfactory recording of impulses in afferent fibres from receptors in the great veins and atria has been accomplished by a number of investigators (Amann & Schaifer, 1943; Walsh, 1947; Whitteridge, 1948; Jarisch & Zotter- man, 1948; Dickinson, 1950; Neil & Zotterman, 1950), and it has been shown that their frequency of discharge is related to the pressure in the great veins and atria (Whitteridge, 1948; Dickinson, 1950). The receptors so far described are known to arise from the atria and are characterized by the presence of an a volley of impulses in time with the a wave of the venous pressure curve. Hereafter these will be referred to as type A atrial receptors. The existence of another kind of cardiovascular fibre in the vagus was reported by Walsh & Whitteridge (1944) and confirmed by Whitteridge (1948). It was shown that this fibre differed in many ways from venous and depressor fibres and, for several reasons, it was believed that it arose from receptors in the small vessels of the lung. Later, Pearce & Whitteridge (1951) showed that a linear relationship existed between the activity ofthese 'pulmonary vascular' fibres and the pulmonary arterial pressure. The blocking temperature of these fibres was shown to be about 8 to 40 C (Torrance & Whitteridge, 1948) which corresponded well with their conduction velocity (Paintal, 1952). However, no experiments with the view to locating these pulmonary vascular receptors had been undertaken with the open chest so far. This was recently achieved, and in this paper the results of these experiments will be described. It will be shown that all such receptors encountered so far arose in the right and left atria of the heart; these will be referred to as type B atrial receptors. METHODS Twelve cats anaesthetized with chloralose (80 mg/kg) were used in the present series of experi- ments; satisfactory results were obtained from ten. The technique of isolating single units and recording their nerve impulses was identical with that described elsewhere (Paintal, 1953). All the experiments were done on the right cervical vagus. The right venous pressure was recorded by a semi-rigid catheter 20 cm long inserted through the left external jugular vein and connected to a capacitance manometer filled with 0-9 % (w/v) * Present address: T. D. E. Laboratory, Kanpur, India. Downloaded from J Physiol (jp.physoc.org) by guest on April 19, 2011 ATRIAL RECEPTORS 597 sodium chloride. In one experiment the right intraventricular preasure was recorded with a similar catheter passed down the left external jugular vein and the catheter recording the venous pressure was inserted down the corresponding vein of the right side. The electronic circuit used was a modification of the one described by Alexander (1951) for an ultramicrometer. Additional amplification was obtained by a direct-coupled pentode which permitted a sensitivity of over 20 V/cm H2O. The manometer was linear over the range of pressures recorded and was stable over long periods. Nevertheless, to obviate any errors arising from base-line shift, each record was calibrated separately with a water manometer. The natural period of the whole recording system filled with salinewas about 50 c/s. The positions of the catheters were always checked at the end of each experiment. A constant endeavour was made to identify artifacts due to movements of the heart in the venous pressure records. Provided that the catheter was not blocked at the time of recording the pressure records were, as a rule, free from artifacts; where they were present the pressure records were discarded. The intrapleural pressure was recorded with a wide bore needle connected to an air-filled mirror membrane manometer. RESULTS Location of so-called pulmonary vascular receptors Altogether, the location of ten such fibres has been achieved with certainty. Experiments were always begun with the animal breathing spontaneously with an intact chest. Single units were dissected until a pulmonary vascular fibre was obtained (Fig. 1) and was then finally identified by the criteria L........... _____ II _ A .1I-IA - Fig. 1. A type B right atrial fibre. From above downwards, e.c.g.; impulses in a right atrial fibre; time in see; intrapleural pressure, inspiration downwards. Activity in the fibres is increasing during inspiration. described already (Whitteridge, 1948; Pearce & Whitteridge, 1951). These were, briefly: (1) that the pattern of discharge should be of a late systolic character, without an a burst of impulses in time with auricular systole; (2) the activity should be augmented by an increase in venous return produced by normal or obstructed inspiration, or by suction of air from the trachea; (3) the activity should be decreased or abolished during a sustained positive pressure inflation of the lungs. Downloaded from J Physiol (jp.physoc.org) by guest on April 19, 2011 598 A, S. PAINTAL Identification of the fibre was followed by putting the animal on positive pressure ventilation and opening the chest as quickly as possible. This was done by cutting through the mid-sternal region, taking care that the nerve fibre on the recording electrodes was not disturbed at all. In order to make the necessary observations as quickly as possible no attention was paid to haemo- stasis as the viability of single unit preparations is very uncertain. Fig.... 2'O. Effect of occluding:the:pulmonaryarteryonthe fibre .........5.......s n Fg.:.n . ... 1. R ds A.B and C arrow in-A and released atarro,w ..in --. type of cardiac rhythm with intervals of silence, despite the contine re 5, ^,;' 5 t .' v ; < - : y: - ^ sr s S.'t.....; 5. 4 _wS°i I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~........~s =_F 2'A~~~~~~~~~~~~~~~....... ...... .@''2...''.''''.' .. .... .. .. .. .. ,. .. .. .. .. .. .. .? j2 ... ... .. .. .. ... .. .. .. .... .... .... ....: ... .. .. .. .. ..-i arcotos. Fro aoe donars right intr.ara pesr; e.cA.g.; imule in a rIghA,.5tatia fibre; time,in + se. Ocl,o of th pumnr arer was comne at andthumb. In six fibres there was an; immediate increase in; fibre discharge "~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~........ .................. .: the~~~~~~~~~~~~~gcoprsso on. th pum nr arer w)A,,smaintie. Th clsion;-S ;-i . ~~~~~~~~~~~. ..... ...... ... ......... ...... .. Fig. 2. Effect of occluding the pulmonary artery on the fibre shown in Fig. 1. Records A, B and C are continuous. From above downwards, right intra-atrial pressure; e.c.g.; impulses in a right atrial fibre; timne in -1- sec. Occlusion of the pulmonary artery was commenced at arrow in A and released at arrow in C. The pericardium was slit and the pulmonary artery occluded between finger and thumb. In six fibres there was an immediate increase in fibre discharge associated with an increase in venous pressure (Fig. 2) which lasted as long as the compression on the pulmonary artery was maintained. The occlusion initially produced a continuous discharge which then regained the original type of cardiac rhythm with intervals of silence, despite the continued rise Downloaded from J Physiol (jp.physoc.org) by guest on April 19, 2011 ATRIAL RECEPTORS 599 in venous pressures. The silent intervals corresponded to the fall in the venous pressure, and they disappeared when the right auriculo-ventricular (A-v) junc- tion was compresed, which produced a much greater rise in venous pressure. The interval of silence in spite of the raised venous pressure is probably due to the sudden decrease in the stimulus intensity-an explanation provided by Bronk & Stella (1934) for the behaviour of carotid pressure receptors. 1 2 EI 24 E4.12 .. 24 -........ I- 12 _- UElOL Fig. 3. Effect of occluding the pulmonary artery on a type B left atrial fibre. A, B and C are continuous. From above downwards, right intra-atrial pressure; right intraventricular pressure; e.c.g.; impulses in fibre; pulmonary artery was occluded between upper and lower arrows. The calibrations apply to the intraventricular pressure. In four fibres occlusion ofthe pulmonary artery abolished the fibre discharge which returned after a variable period on releasing the occlusion. This is illustrated in Fig. 3 which shows the effectiveness of the occlusion by the rise in the right intraventricular pressure. In fibres responding in this way compression of the left A-v junction produced a pronounced increase in fibre activity which always lost its cardiac rhythm and became continuous (Fig. 4). On releasing the occlusions the fibre discharge regained its original character. Compression ofthe left A-v junction produces a rise in the right intraventricular and the right venous presures as well, although the rise was never as great as that produced by occluding the pulmonary artery. For this reason occlusion of Downloaded from J Physiol (jp.physoc.org) by guest on April 19, 2011 600 A. S. PAINTAL IMAPShhhaAAwAw Fig. 4. Effect of occluding the left A-V junction on a type B left atrial fibre. The records are continuous. From above downwards, e.c.g.; impulses in the fibre; time in tl- sec. Left A-V junction was occluded between arrows. U Fig. 5. Effect of clamping the pulmonary veins on two type B left atrial fibres. A and B are continuous. From above downwards, right intraventricular pressure (calibrated); right intra-atrial pressure; e.c.g.; impulses in fibres. The clamp on the pulmonary veins was released at arrow in A and re-applied at arrow in B. Note that, at upper arrow, activity in the fibre with a small impulse ceases and activity in the fibre with large impuls begins. The reverse takes place at the arrow in B. This is presumably due to the two fibres being proximal and distal to the clamp respectively. Downloaded from J Physiol (jp.physoc.org) by guest on April 19, 2011 ATRIAL RECEPTORS 601 the left A-V junction also gives rise to increased fibre activity in right atrial fibres.
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